鄱阳湖沉水植物区不同深度土壤甲烷排放对温度水分变化的响应

采集鄱阳湖沉水植物区0-10 cm和10-30 cm土壤样品,通过设置2个温度（18℃和28℃）和2个水分（淹水2 cm和土柱取出水面后的实际土壤水分含量）处理组合,进行持续2年的甲烷（CH₄）排放室内培养实验,以探讨不同深度土壤CH₄排放对温度、水分变化的响应差异,以及温度、水分和土层对湿地土壤CH₄排放的交互影响。结果表明：0-10 cm和10-30 cm土壤CH₄排放速率变化范围分别为0.01-3.63 μgCH₄-C kg^-1d^-1、0.02-1.99 μgCH₄-C kg^-1d^-1;均值分别为0.72和0.15 μgCH₄-C kg^-1d^-1。温度、水分和土层3因素及其交互作用均对土壤CH₄排放有显著影响（P<0.01）,且土层的影响最大。两水分处理下的CH₄排放对温度变化的敏感性均表现为0-10 cm（Q₁₀为1.78、3.26）高于10-30 cm土层（Q₁₀为1.04、1.08）。CH₄平均排放速率及累计排放量均表现为0-10 cm显著高于10-30 cm土层,且培养前期高于培养后期,显示基质有效性对土壤CH₄排放的重要影响。     

潮汐作用对黄河三角洲盐沼湿地甲烷排放的影响

盐沼湿地作为陆海交互作用的过渡带是CH₄重要的自然来源。潮汐活动通过影响CH₄的产生、氧化和传输驱动了湿地CH₄间歇性、周期性的排放。利用涡度相关和微气象监测技术,对黄河三角洲一个盐地碱蓬生态系统CH₄通量、环境因子和水文要素（潮汐）进行了长期连续监测分析了该生态系统生长季CH₄排放的季节动态及潮汐作用对CH₄排放的影响。结果表明：生长季该生态系统是CH₄的排放源,排放日均值为0.063 mg m^-2 h^-1,（范围为-0.36-0.57 mg m^-2 h^-1）。潮汐淹水阶段和落潮后湿润阶段表现为CH₄的显著源。此外我们发现,短期潮汐活动引起土壤干湿状况的变化促进了CH₄脉冲式的排放,因此未来气候变化下温度升高和降雨季节分配引起的土壤干湿变化将会对该区域CH₄排放甚至碳循环产生积极影响。     

鄱阳湖苔草湿地甲烷释放特征

2009年5月-2010年4月在鄱阳湖南矶湿地国家级自然保护区选择以灰化苔草为建群种的洲滩,设置土壤-植物系统(TC)、剪除植物地上部分 (TJ)2个试验处理,利用密闭箱-气相色谱法测定了鄱阳湖典型苔草湿地的甲烷(CH₄)释放通量。结果表明:1)TC、TJ 2个试验处理CH₄释放速率变化范围分别为-0.094-17.75 mg · m^-2 · h^-1、-0.122-19.16 mg · m^-2 · h^-1,均表现出明显的季节变化规律;2)地表未淹水期间,剪草处理CH₄释放显著高于非剪草处理(t=2.69, P<0.05);地表淹水达到15 cm后,剪草处理CH₄释放明显低于非剪草处理。3)土壤5 cm温度、土壤水分与2处理非淹水期间CH₄释放速率均呈显著正相关,是非淹水期间CH₄通量变化的主要控制因子,2因子能够共同解释非淹水期苔草湿地65%-74%的CH₄通量变异;4)试验期间,苔草湿地CH₄释放量约为12.77 gC/m²,相当于同期土壤有机质分解碳排放量的4%,甲烷释放的碳消耗不足苔草湿地年NPP的1%。     

微生物胞外长距离电子传递网络研究进展

[目的] 解析一株从黄河三角洲湿地甲烷氧化富集物中分离获得的甲烷氧化菌伴生菌的生理学及电化学特性,并探究该菌株对甲烷氧化过程的影响。[方法] 使用高通量测序技术解析甲烷氧化富集物的菌群结构,采用稀释涂布法、平板划线法分离甲烷氧化菌的伴生菌,通过16S rRNA基因测序技术进行菌株初步鉴定。利用扫描电子显微镜观察菌株形态,并通过气相色谱（gas chromatography,GC）检测伴生菌利用甲烷情况及对甲烷氧化菌氧化甲烷效率的影响。采用双室微生物燃料电池（microbial fuel cells,MFCs）及差分脉冲伏安法（differential pulse voltammetry,DPV）检测菌株的电化学活性。[结果] 黄河三角洲湿地土壤甲烷氧化富集物主要的好氧甲烷氧化菌为甲基杆菌属Methylobacter,同时还发现一些伴生菌。分离得到一株甲醇利用菌P7,其16S rRNA基因序列与恶臭假单胞菌Pseudomonasputida的相似性达99.79%。扫描电镜结果显示该菌株为杆状,长约1.5-2.5μm,宽度约为0.5μm。GC检测结果显示,该菌株不能利用甲烷,但与甲烷氧化菌共培养时,可以促进甲烷氧化（P<0.05）。双室MFCs检测结果显示该菌株具有电活性,最大电流输出密度为28 mA/m²,DPV检测结果显示该菌株主要的氧化峰和还原峰分别位于-0.17 V和-0.25 V。[结论] 本研究从黄河三角洲湿地甲烷氧化富集物中获得一株具有电活性的甲烷氧化菌的伴生菌恶臭假单胞菌Pseudomonas putida P7,该菌株可以促进甲烷氧化。本研究加深了对甲烷氧化过程中伴生菌的生理学特性及功能的认识。     

贵州阿哈湖水库沉积物中甲烷氧化菌分布与功能

甲烷氧化菌是一类能够以甲烷作为唯一碳源和能量维持生存的微生物,其活动与生态系统中物质循环及能量流动密切相关。【目的】了解阿哈湖水库沉积物中甲烷氧化菌群落结构及代谢功能。【方法】采用宏基因组技术对环湖沉积物和湖心沉积物进行研究。【结果】水库沉积物中主要的好氧甲烷氧化菌是甲基杆菌属(Methylobacter) (0.37%)和甲基单胞菌属(Methylomonas) (0.12%),主要的厌氧甲烷氧化菌是Candidatus_Methylomirabilis (0.12%),属于NC10门细菌中的亚硝酸盐反硝化型厌氧甲烷氧化菌,其中好氧甲烷氧化菌的pmoA基因为6.16×10⁷ copies/g,反硝化型厌氧甲烷氧化菌的16S rRNA基因为2.84×10⁷ copies/g。4种代谢的功能基因多样性表现为氮代谢>碳代谢>硫代谢>甲烷代谢,基于京都基因和基因组百科全书(Kyoto encyclopedia of genes and genomes, KEGG)基因库进行注释得到6大类功能,发现18条与碳(包括甲烷)、氮、硫代谢有关的完整代谢途径。主坐标分析(principal coordinates analysis, PCoA)显示环湖沉积物与湖心沉积物之间的甲烷氧化菌种类和功能存在显著差异。影响甲烷氧化菌分布的主要环境因子为氧化还原电位、电导率和硫酸根。【结论】阿哈湖水库甲烷氧化菌以Ⅰ型好氧甲烷氧化菌为主,甲烷氧化菌群落代谢途径丰富,Ⅰ型和Ⅱ型甲烷氧化菌在对O₂的适应性上有显著差异。相关研究可为湖泊水生态环境的保护和微生物资源的开发利用等方面提供一定的理论支撑。     

典型温带雨养泥炭沼泽湿地地下部二氧化碳和甲烷浓度变化规律及其影响因素

为研究北方泥炭沼泽湿地二氧化碳（CO₂）和甲烷（CH₄）浓度随深度的变化规律及其影响因素,选取欧洲北部典型雨养泥炭地贝尔山湿地（BBM）和舒特兹山湿地（SBM）两个采样点,通过原位采集泥炭剖面温室气体、孔隙水以及土壤样品,结合傅里叶变换红外光谱（FTIR）技术、碳氮同位素技术,探讨泥炭土壤的分解程度及温室气体浓度变化的关系。研究结果表明：（1）BBM采样点地下部的CO₂浓度变化规律总体呈现随深度波动减少趋势,值多在3000 μmol/L附近波动,最大值为4210.74 μmol/L（120 cm）,SBM采样点的CO₂浓度随深度先增后减,60 cm以上在1800 μmol/L附近波动,60 cm以下在3000 μmol/L附近波动,最大值为4191.94 μmol/L（90 cm）;BBM和SBM地下部CH₄浓度都随深度增大,并且在60cm以下浓度增加较快,BBM最大值为735.90 μmol/L（260 cm）,SBM最大值为543.51 μmol/L（170 cm）。（2）BBM和SBM的δ¹³C_CH4的值均较小,表明大部分的¹²CH₄仍被储存在泥炭土壤中,而两个采样点的δ¹³C_CO2和分馏系数α_c均随深度增加,表明泥炭土中产甲烷方式为浅层以乙酸产甲烷为主,深层以H₂还原CO₂为主。（3）C/N、碳氮同位素比值、FTIR均显示SBM和BBM的有机质分解程度较低,因为两个采样点的低可溶性有机碳浓度和低pH值不利于分解,表明该地储存着大量有机碳。通过探讨北温带典型泥炭地温室气体的浓度变化规律及其影响因素,结果可为全球泥炭湿地碳排放提供理论支撑。     

若尔盖高原沼泽湿地CO2排放时空变化特征

为了更好理解若尔盖高原不同微生境下沼泽湿地生态系统CO₂排放通量的变化特征,以若尔盖高原湿地自然保护区为研究对象,2013和2014年生长季期间,采用了静态箱和快速温室气体法原位观测了3种湿地5种微生境下沼泽湿地CO₂排放通量时空变化规律。结果表明：长期淹水微地貌草丘区湿地（PHK）和洼地区湿地（PHW） CO₂排放通量变化范围分别为38.99-1731.74 mg m^-2 h^-1和46.69-335.22 mg m^-2 h^-1,季节性淹水区微地貌草丘区湿地（SHK）和洼地区湿地（SHW） CO₂排放通量变化范围分别为193.90-2575.60 mg m^-2 h^-1和49.93-1467.45 mg m^-2 h^-1,而两者过渡区的无淹水区沼泽湿地（Lawn） CO₂排放通量变化范围194.20-898.75 mg m^-2 h^-1。相关性分析表明5种微地貌区沼泽湿地CO₂排放通量季节性变化与不同深度土壤温度均存在显著正相关,与水位存在显著负相关（PHW、SHW、SHK、Lawn）或不相关（PHK）,并且水位和温度（5 cm）共同解释了CO₂排放通量季节性变化的87%。3种湿地5种微生境下沼泽湿地CO₂排放通量存在空间变化规律,主要受水位影响,但植物也影响沼泽湿地CO₂排放通量空间变化规律,并且表明沼泽湿地CO₂排放通量与水位平均值存在显著负相关。     

内陆湿地与水体甲烷厌氧氧化功能微生物研究进展

内陆湿地与水体(如湖泊、河流、水库等)是温室气体甲烷的重要排放源。微生物介导的甲烷厌氧氧化(anaerobic oxidation of methane,AOM)反应在控制内陆湿地与水体甲烷排放中起着不可忽视的作用,对缓解全球温室效应具有重要意义。内陆湿地与水体易形成缺氧环境,且电子受体的种类和数量繁多,是发生AOM反应的理想生境。近年来,不断有研究表明,内陆湿地与水体中存在多种电子受体(NO₂^-、NO₃^-、SO₄^2-、Fe (III)等)驱动的AOM途径。NC10门细菌和甲烷厌氧氧化古菌(anaerobic methanotrophic archaea,ANME)的一新分支ANME-2d主导了湿地和水体环境中的AOM反应,其中ANME-2d具有根据环境条件选择不同电子受体的潜力。研究系统综述了内陆湿地与水体中不同电子受体驱动的AOM途径及其参与的主要功能微生物类群;分析了AOM反应在控制温室气体甲烷排放中的作用及其环境影响因素;总结了相关功能微生物的分子生物学检测方法及甲烷厌氧氧化活性测定的同位素示踪技术。最后,对未来相关研究方向进行了展望。     

典型油气藏区域环境因素差异对甲烷氧化菌丰度与群落结构分布的影响

【目的】甲烷氧化菌(methane-oxidizing bacteria, MOB)是油气微生物勘探中重要的指标微生物,其丰度与群落结构分布受到地理位置和多种环境因素的影响。本研究以采集的7个典型油气藏区域的土壤样品为研究对象,探究油气藏区域环境因素差异对甲烷氧化菌丰度与群落结构分布的影响。【方法】对采集自7个典型油气藏区域土壤样品进行pmoA基因实时荧光定量PCR (real-time fluorescence quantitative PCR, qPCR)、细菌16S rRNA基因测序和甲烷氧化菌功能基因pmoA测序,结合环境因子,比较甲烷氧化菌丰度差异,分析环境因子对甲烷氧化菌分布的影响。【结果】土壤样品的理化性质测定发现江汉盆地样品的含水量最高,约22.8%,硝态氮平均含量最高位于玉北油田,达到31.96 μg/g干重土壤,春光油田出现最高的SO₄^2- (6 425.0 mg/g干重土壤)及Cl^- (1 617.0 mg/g干重土壤)浓度。qPCR分析发现pmoA基因丰度仅为土壤样本总细菌丰度的0.77%,表明甲烷氧化菌绝对数量较少。16S rRNA基因测序发现type I型甲基八叠球菌属(Methylosarcina)、甲基嗜热菌属(Methylocaldum)、甲基球菌属(Methylococcus)与type II型的甲基胞囊菌属(Methylocystis) 4种主要的甲烷氧化菌,甲烷氧化菌中的优势菌属的相对丰度极低,最高仅为0.124%。甲烷氧化菌功能基因pmoA测序发现type II型的甲基胞囊菌属、甲基弯曲菌属(Methylosinus)为主要优势pmoA基因型。环境因子与甲烷氧化菌丰度相关性分析表明甲烷氧化菌绝对丰度与铵态氮、pH、颗粒大小、硫酸根和氯离子显著相关,而环境因子与甲烷氧化菌的相对丰度的相关性分析仅发现与颗粒大小、总氮、总磷、金属离子(Al、Fe、K、Ca、Mg、Mn、Zn和Cu)显著相关。甲烷氧化菌pmoA功能基因组成判别相关分析(discriminant correlation analysis, DCA)表明含水量(土壤湿度)、pH值、<2 μm的土壤颗粒、总氮及钙离子对甲烷氧化菌pmoA功能基因组成影响具有显著性。【结论】本研究通过对来自7个典型油气藏区域的土壤样品的甲烷氧化菌的绝对丰度与相对丰度进行分析,发现基于不同的测量方法,甲烷氧化菌的丰度存在较大差异,16S rRNA基因测序无法较为全面地反映土壤样本中优势甲烷氧化菌的群落结构,不同地理位置的甲烷氧化菌指示群落存在差异,在全国尺度下油气区及背景点均无特有的甲烷氧化菌指示群落。对环境因子与甲烷氧化菌相关性分析发现,基于不同的甲烷氧化菌丰度指标,环境因子的显著影响具有较大差别。     

太湖流域农田稻季CH4通量特征及影响因子

开展太湖流域农田稻季CH₄排放研究,深入了解稻田CH₄排放规律,为稻田CH₄减排、制定合理稻田管理措施提供科学依据。以太湖流域稻麦轮作农田为研究区域,运用涡度相关法观测其稻季CH₄通量变化,分析其通量变化特征及影响因子。结果表明：太湖流域典型稻麦轮作区稻季为CH₄的源,CH₄排放总量为28.95 g/m²,稻季CH₄通量日变化表现为无规则型与单峰型两种模式;稻季CH₄排放整体集中在水稻生长前期（81.61%）及中期（16.16%）、后期排放相对较弱（2.23%）,返青期排放量较低（日均0.102 μmol m^-2 s^-1）,分蘖期较强（日均0.451 μmol m^-2 s^-1）,成熟期最低（日均0.006 μmol m^-2 s^-1）;模型所模拟的累计CH₄排放通量比累计测量CH₄通量低6.69%,较好地模拟了太湖流域稻田CH₄的排放,土壤温度、土壤水分、土壤电导率、摩擦风速可确认为太湖流域农田稻季CH₄排放的主要驱动因子。     

Succession in the southern part of the Canadian boreal forest

Forest succession following fire in a forest mosaic of northwestern Quebec has been studied in order to: (1) describe the successional pathways using communities of different ages and (2) evaluate convergence of successional pathways and possible effect of fire suppression on the establishment of steady-state communities. As a first step, ordination and classification techniques were used in order to remove changes in forest composition which are related to abiotic conditions. Then, ordinations based on tree diameter distributions were used to study shifts in species composition in relation to time since the last fire.Even under similar abiotic conditions, successional pathways are numerous. However, regardless of forest composition after fire, most stands show convergence toward dominance of Thuja occidentalis and Picea mariana on xeric sites and dominance of Abies balsamea and Thuja occidentalis on more mesic sites. Stable communities of >300 yr occur on xeric sites while on mesic sites directional succession still occurs after 224 yr. Nearly all species involved in succession are present in the first 50 yr following fire. Only Abies balsamea and Thuja occidentalis increase significantly in frequency during succession. Following initial establishment, successional processes can generally be explained by species longevity and shade tolerance. Early successional species may be abundant in the canopy for more than 200 yr while the rapid decrease of Picea glauca, a late successional species could be related to spruce budworm outbreaks. Considering the short fire rotation observed (about 150 yr), a steady-state forest is unlikely to occur under natural conditions, though it may be possible if fire is controlled.     

Vermicomposting in the management of pig-waste in Hong Kong

The treatment and disposal of pig-waste in Hong Kong has received much attention in recent years but, following any of the presently used treatment processes, solids remain to be further stabilized. Vermicomposting is a waste stabilization technique which converts waste into potentially recyclable materials such as worm protein and worm casts. The earthworm, Pheretima asiatica, can stabilize most of the solids arising from the treatment of pig-waste, including raw pig manure, suggesting that vermicomposting has a high potential as a unit process in the management of pig-waste in Hong Kong.S.H. Wong is with the Environmental Protection Department, Hong Kong; and D.A. Griffiths is with the Department of Botany, University of Hong Kong, Hong Kong.     

Research on streptococci in the UK

Pleurotus florida produced high amounts of laccase (4.60 U/ml) in malt extract broth after 12 days' growth under stationary conditions. The production of laccase was semi-constitutive. Hyperlaccase mutants ofP. florida were obtained through mutagenesis of mycelial protoplasts usingN-methyl-N-nitro-N-nitrosoguanidine (50 g/ml) for 2 min. Three hyperlaccase mutants were selected showing growth and enzyme production responses similar to the parent.     

New and old problems in the taxonomy of the Gelidiales (Rhodophyta)

The order Gelidiales includes over 140 agar-producing taxa. Many species are taxonomically confused; the boundaries of one family and all but four genera recently have been contested, and the controversy over ordinal status has lasted for over 25 years. This study reviews recent developments in the taxonomy of the group and suggests future areas for studies. The order cannot be defined by a few exclusive characters, as intended in the past, but it shows a unique combination of characters. Additional studies on Gelidiella and Acanthopeltis seem advisable to clarify family limits. The segregation of Onikusa and Pterocladiastrum is doubtful. Only three of six characters discriminate Gelidium from Pterocladia. None allows complete generic segregation and all need variability studies. Analysis of nomenclatural types in the light of morphological variation would permit an understanding of species limits in Gelidium and Pterocladia. Similar studies are needed in Gelidiella and Ptilophora.     

Intralineage variation in the pattern ofrbcL nucleotide substitution

Variation in chloroplastrbcL sequences was studied in representative species of four different lineages: the tribeRubieae (Rubiaceae), and the generaDrosera (Droseraceae),Nothofagus (Nothofagaceae) andIlex (Aquifoliaceae). Each lineage has its particular non-overlapping set ofrbcL polymorphic sites, indicating that common unconstrainedrbcL sites are not shared. Large differences in the rate and pattern of nucleotide substitution are observed among the four lineages. The genusIlex has the lowest rate of substitution, the lowest transition/transversion ratio, the lowest synonymous/replacement ratio and the lowest number of substitutions at the third codon position. An apparent relationship of these measures to the age of the lineages is observed. The A + T content and codon use among the four lineages are very similar and, apparently, cannot account for the observed differences in patterns of nucleotide substitution. However, the A + T content of the two bases immediately flanking the polymorphic sites is higher inIlex than in the other lineages. This could be correlated with the transversion/transition bias observed inIlex. The particularly low synonymous/replacement ratio found inIlex could also be explained by the small population sizes of species in this genus.     

Patterns in the distribution of myctophid fish in the northern Scotia Sea ecosystem

The mesopelagic fish community of the northern Scotia Sea was investigated during the austral autumn using multi-frequency acoustics, opening and closing nets and pelagic trawls fished from the surface to 1,000 m. The Family Myctophidae (15 species in 5 genera) dominated the ichthyofauna, with larval notothenids caught over the South Georgia shelf and bathylagids and stomiids abundant in deeper hauls. The biomass of myctophids was estimated to be 2.93 g wet weight 1,000 m⁻³, with Electrona carlsbergi, E. antarctica, Protomyctophum bolini, P. choriodon, Gymnoscopelus braueri, G. fraseri, G. nicholsi and Krefftichthys anderssoni, being the most abundant species. Analysis of community structure indicated a high level of depth stratification within the myctophids, with evidence of diurnal vertical migration in some, but not all, species. Length-frequencies of G. braueri, G. nicholsi, E. antarctica and K. anderssoni were multimodal, suggesting that all life stages may be present in the northern Scotia Sea. In contrast, P. choriodon, P. bolini, G. fraseri and E. carlsbergi had unimodal distributions despite having multi-year lifecycles, indicating that they probably migrate into the region from warmer areas to the north.     

Fungal biodiversity in the air of Turin

The qualitative fungal composition of Turin 's atmospheric environment was surveyed, carrying out a twelve-month study and collecting with a single stage volumetric sieve sampler on Dermasel agar supplemented with 0.4 g l^–1 cycloheximide and 0.05 g l^–1 chloramphenicol. We isolated 165 species and 2 varieties of mesophilic fungi from 58 genera and 26 thermotolerant species from 12 genera. Penicillium, Aspergillus, Acremonium, Chrysosporium, Scopulariopsis, Malbranchea, Paecilomyces, Phialophora and Cladosporium were in sequence the genera most rich in mesophilic species; Aspergillus, Penicillium, Chrysosporium and Scopulariopsis the most rich in thermotolerant species. Many of the species isolated are rarely or never recorded in the atmospheric environment. Cycloheximide can thus be said to select among airborne fungi, giving a characteristic picture.Abbreviation CH Cycloheximide - cfu colony forming unit     

New Discoveries in the Canellaceae in the Antilles: How Phylogeny can Support Taxonomy

Six genera have been described in the family Canellaceae, four of them from the Neotropics and the other two from Africa and Madagascar. The Caribbean genera are Canella, Pleodendron and Cinnamodendron. Canella is a monotypic genus widespread in the region, and Pleodendron is present in the Greater Antilles and Costa Rica. Cinnamodendron occurs in the Greater Antilles (Cuba, Hispaniola, and Jamaica) as well as in South America. A recent phylogenetic analysis of the family shows that Cinnamodendron is not monophyletic because the South American species and the Antillean species are recovered in two different clades. The Antillean species formed a clade sister to Pleodendron. The synapomorphies of the Antillean species of Cinnamodendron are tetramerous flowers with eight petals, eight stamens, four carpels, and four placentae. Based on the results from the phylogenetic analysis major taxonomy changes are expected for the family.

---

Resumen  Seis géneros han sido descritos en la familia Canellaceae, cuatro de estos para el neotrópico y los otros dos para Africa y Madagascar. En las Antillas se encuentran los géneros Canella, Pleodendron y Cinamodendron. Canella es un género monotípico de amplia distribución en la región del caribe y pleodendron se encuentra presente sólo en las antillas mayores y Costa Rica. Cinnamodendron ha sido dado tanto para las Antillas mayores como para America del Sur. Un análisis filogenético previo de la familia indica que el género Cinnamodendron no es monofilético. Existe una separación de las especies Sudamericanas y Antillanas en clados diferentes. Las especies de las Antillas forman un clado que es hermano de Pleodendron. Los carácteres sinapomórficos de las especies antillanas de Cinnamodendron son: flores tetrámeras con ocho petalos, ocho estambres, cuatro carpelos y cuatro placentas. Basados en los resultados de la filogenia del grupo, se anticipan cambios taxonómicos para la familia.

     

Hybridization and speciation in the genusDryopteris (Pteridophyta:Dryopteridaceae) on Pico Island in the Azores

The cytology of wild hybrids ofDryopteris from Pico, Azores has been examined. This has confirmed the parentage of the hybrids, and provided evidence to support the view that the tetraploid speciesD. crispifolia has evolved in the Azores from two diploid species,D. aemula andD. azorica, by hybridization and polyploidy.